Registration of 'Cara' Soft White Winter Club Wheat

c u lt i v a r

Journal of Plant Registrations, Vol. 7, No. 1 81

Registration of ‘Cara’ Soft White Winter Club WheatKim Garland-Campbell,* Robert E. Allan, James Anderson, Adrienne Burke, Nathan Blake, Chris Hoagland, Carl Walker, Jeron Chatelain, Lynn M. Little, John Pritchett, Xianming Chen, Craig Morris, Deven See, Stephen Guy, Timothy Murray, Doug Engle, Henry Wetzel, and Dave Wood

ABSTRACTClub wheat (Triticum aestivum ssp. compactum) is an important component of the export grain market for the Pacific Northwest. Our objective was to develop a club wheat cultivar with resistance to stripe rust (caused by Puccinia striiformis f. sp. tritici Westend.) and strawbreaker foot rot [caused by Oculimacula yallundae Crous & W. Gams (Wallwork & Spooner) and O. acuformis (Boerema, R. Pieters & Hamers) Crous. & W. Gams] and with the end-use qualities that make club wheat a specialty product. The bulk pedigree breeding method was used to select ‘Cara’ (Reg. No. CV-1078, PI 643435) from the cross WA7752//WA6581/WA7217 made in 1992. Cara is a semidwarf wheat with the Rht-D1b dwarfing allele. Cara has winter hardiness equal to that of other club wheat cultivars—Bruehl, Chukar, and Coda—but less than the soft white wheat cultivars Eltan (PI 536994) or Masami (PI 634715). Cara was evaluated in multi-environment replicated plot trials in the Washington since 2003. Grain yields were equal to or better than other club and soft white cultivars in the 30- to 40-cm and the 40- to 50-cm annual precipitation zones. The milling and baking quality of Cara was equal to those of the best club wheat check cultivars. Cara is resistant to all races of stripe rust prevalent since 2001, possessing Yr17 and unknown genes for stripe rust resistance. It is also resistant to strawbreaker foot rot, possessing Pch1. Cara provides growers with an agronomically competitive cultivar with resistance to stripe rust and strawbreaker foot rot plus excellent club wheat end-use quality.

K. Garland-Campbell, A. Burke, X. Chen, C. Morris, D. See, and D. Engle, USDA-ARS, Wheat Genetics, Quality, Physiology and Disease Research Unit, 209 Johnson Hall, Washington State Univ., Pullman WA 99164-6420; R.E. Allan, L.M. Little, J. Pritchett, and D. Wood (all retired), USDA-ARS, Wheat Genetics, Quality, Physiology and Disease Research Unit, 209 Johnson Hall, Washington State Univ., Pullman WA 99164-6420; J. Anderson, Dep. of Agronomy and Plant Genetics, Univ. of Minnesota, 411 Borlaug Hall, 1991 Upper Buford Cir., St. Paul, MN 55108-6026; N. Blake, Connell Grain Growers, 3132 Road O NE, Moses Lake, WA 98837; C. Hoagland, Dep. of Agron-omy, Purdue Univ., 915 West State St., West Lafayette, IN 47907-2054; C. Walker and S. Guy, Dep. of Crop and Soil Sciences, Box 646420, Washington State Univ., Pullman, WA 99164-6420; J. Chat-elain, Oregon State Univ., Columbia Basin Agricultural Research Center, 48037 Tubbs Ranch Rd., P.O. Box 370, Pendleton, OR 97801; T. Murray and H. Wetzel, Dep. of Plant Pathology, Box 646430, Washington State Univ., Pullman, WA 99164-6430. *Corresponding author ([email protected]).

Abbreviations: PNW, Pacific Northwest; WSU, Washington State Univ.

Club wheat (Triticum aestivum ssp. compactum) com-prises 5 to 10% of the winter wheat crop in the Pacific

Northwest (PNW) states of Washington, Oregon, and Idaho (USDA-NASS, 2011). Club wheat has a unique spike mor-phology, conditioned by the dominant compactum (C) locus on chromosome 2DL (Johnson et al., 2008), that results in redistribution of yield components. Club cultivars tend to have smaller seeds than lax cultivars. Although the rachis length and spike are shorter than in lax genotypes, the spikelet fertility and number of seeds per spike is greater in club wheat (Zwer et al., 1995). Club wheat is marketed as a value-added product to the countries in the Asian rim as a component of the Western White wheat market class (a blend of the soft white wheat subclasses soft white wheat plus 10 to 20% white club wheat [USDA, 2006]). Western White is valued for its consistent excellent cake- and cookie-baking qualities and frequently commands a premium over other soft wheat market classes. Because of the established market based on end-use quality, club wheat cultivars have consistently been bred to have greater flour extraction and break flour yields, higher starch content, and greater stabil-ity for low starch damage than soft white winter wheat (Lin and Czuchajowska, 1997; Morris et al., 2005). Flour milled from club wheat has the low viscosity and weak gluten quality that is desired for air-leavened cake products.

Club wheat acreage in Washington and Oregon state is largely planted to two cultivars developed by Washington State University (WSU) and the USDA-ARS: Bruehl

Published in the Journal of Plant Registrations. doi: 10.3198/jpr2012.01.0005crcReceived 5 Jan. 2012. Registration by CSSA. © Crop Science Society of America 5585 Guilford Rd., Madison, WI 53711 USAAll rights reserved. No part of this periodical may be reproduced or trans mitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permission for printing and for reprinting the material contained herein has been obtained by the publisher.

c u l t i v a r82 Journal of Plant Registrations, Vol. 7, No. 1

(PI 606764, Jones et al., 2001) and Chukar (PI 628641, Garland-Campbell et al., 2005). Small amounts of three other cultivars—Coda (PI 594372, Allan et al., 2000), Edwin (PI 606765, Jones et al., 2000), and Moro (CItr 13740, Rohde, 1966)—are also in production. Spring club cultivars include Eden (PI 630983) and JD (PI 656790, Kidwell et al., 2004). Bruehl, Chukar, Coda, and JD club wheat cultivars possess resistance to stripe rust (caused by Puccinia striiformis f. sp. tritici Westend), which is a major disease limiting grain yields in the PNW. Bruehl is susceptible to strawbreaker foot rot [caused by Oculimacula yallundae Crous & W. Gams (Wallwork & Spooner), syn. Pseudocercosporella herpotrichoides (Fron) Deighton var. herpotrichoides; and O. acuformis (Boerema, R. Pieters & Hamers) Crous. & W. Gams, syn. Pseudoecercosporella herpotrichoides var. acuformis Nirenberg].

Bruehl, Chukar, and Coda are tall relative to the average height of other winter wheat cultivars grown in the PNW. Additional cultivar choice would increase growers’ ability to incorporate genetic diversity into their management plans. Our objective was to develop a club wheat cultivar possessing resistance to stripe rust and strawbreaker foot rot, excellent straw strength, and the milling and baking properties that make club wheat a specialty product.

‘Cara’ (Reg. No. CV-1078, PI 643435) club wheat was developed by the USDA-ARS with assistance from the Washington Agricultural Experiment Station and the Oregon Agricultural Experiment Station. Cara was tested under experimental numbers ARS97135 and ARS97135-9 and is derived from a population with the cross number 92X102. It was released in February 2007 because of its combination of yield potential and the quality characteristics desired for the club wheat market class, plus resistance to multiple diseases, including stripe rust, powdery mildew (caused by Erysiphe graminis DC. f. sp. tritici Em. Marchal), and strawbreaker foot rot. Cara is derived from a combination of Clarence J. Peterson and Robert E. Allan’s germplasm and was released because its diverse pedigree broadened the genetic base of the club wheat cultivars currently in production. The recommended area of adaptation for Cara is the intermediate to high rainfall regions of the Palouse (380–600 mm annual precipitation, primarily in winter), where foot rot, stripe rust, and powdery mildew are problems.

MethodsThe pedigree of Cara is WA7752//WA6581/WA7217. The cross was made by R. E. Allan in 1992. The pedigree of WA7752 (sister line to Coda) is ‘Tres’ (CItr 17917)/‘Madsen’ (PI 511673)//Tres (Allan et al., 1986, 1989, 2000). The pedi-gree of WA6581 (PI 486428) is ‘Omar’ (CItr 13072)/Vogel selection 3 (1834, CItr 12692)//P 178383/Vogel selection 1 (CItr 13431)/3/Vogel selection 4 (CItr 13645)/‘Gaines’ (CItr 13448). PI 178383 was collected by J. Harlan in Hakkari, Turkey, and was widely used in the PNW as a source of resistance to bunt [caused by Tilletia caries (DC.) Tul. & C. Tul., syn. T. tritici (Bjerk.) G. Winter and T. foetida (Wallr.) Liro, syn. T. laevis Kühn], and stripe rust. The pedigree of WA7217 (PI 561035) is ‘VPM’ (PI 519303)/ ‘Moisson’ (PI 315998) #951/2* ‘Barbee’ (CItr 17417, Peterson et al., 1977).

The F1 was advanced in the field without selection. The F2 bulk population was planted in a 3.75-m2 plot in the field at the WSU Spillman Agronomy Farm, Pullman, WA in the fall of 1993. In the summer of 1994, 60 heads were collected without selection from the F2 and planted as F3 headrows, 1 m long, four rows per range, 0.3 m apart with 0.1 m between rows at Spillman Farm. In 1995, those headrows were selected visually for height and maturity relative to the commercial check cultivars Madsen, Hiller (PI 587026, Peterson et al., 1999), and Coda, which were planted at intervals within the headrow plots. The F3 headrows were also selected for resistance to stripe rust (see below for stripe rust inoculation methods). Selected headrows were harvested using hand sickles, threshed using a Vogel thresher, and planted in unreplicated F4 plots 3.75 m2 at the Spillman Farm in the fall of 1995. The 3.75-m2 plot size was maintained for all further testing. Selection in the F4 generation was based on agronomic characteristics (heading date, height, standability), disease resistance, grain yield, and test weight.

Heading date was recorded as days from 1 January to 50% of full spike emergence from the boot (Zadoks growth stage 60) on a plot basis (Zadoks et al., 1974). Height was measured after senescence and recorded as the average distance from the base of the plant to the top of the spike on a plot basis. Grain was harvested with a Wintersteiger Classic small plot combine (Wintersteiger) and grain yield was measured on a plot basis and converted to kilograms per hectare. Test weight was recorded based on a cleaned, 600-g sample using AACC International approved method 55-10.01 (2008).

The selected line, ARS97-135 was advanced to the ARS preliminary nursery with three replications at the Spillman Farm in 1996. The preliminary nursery was evaluated for all of the above traits, plus resistance to strawbreaker foot rot and end-use quality characteristics (for methods, see below). Evaluation for all traits was continued for several more generations due to personnel changes in the breeding program. ARS97-135 was evaluated in the ARS advanced trials at three locations (Spillman Farm, WA; Walla Walla, WA; and Lind, WA) from 1997 to 2000 with three replications at each location. Purification was initiated in the F6 generation in 1997, when 40 F7 heads were collected from a bulk increase of ARS97-135 that was planted at Central Ferry, WA. Those headrows were planted as above at Spillman Farm and evaluated for uniformity in 1998. Ten of those headrows were selected, harvested, and evaluated in the ARS purification trial at Spillman in 1999. The selected F7:9 purified line, ARS97-135-9, was used as a seed source to replace ARS97-135 in all trials after 2000 and renamed Cara in 2007. Cara was evaluated in the ARS elite trials since 2001 at Genesee, ID; Hermiston, Lexington, Pendleton, and Moro, OR; and Harrington, Lind, Pullman, and St. Andrews, WA with three replications at each location. Cara was evaluated in the Western Regional Soft Winter Wheat Nursery at 13 locations in Oregon, Idaho, Montana, and Washington in 2003. Comparisons between Cara and check cultivars grown in the 2007–2009 ARS elite trials were made using t tests for independent samples with the

c u l t i v a rJournal of Plant Registrations, Vol. 7, No. 1 83

Satterthwaite approximation based on data from 2007 to 2009 (11–29 location-years, depending on the comparison).

Cara has been evaluated in state extension variety trials in Oregon since 2006, Idaho since 2006, and Washington since 2004. Specifics of plot sizes, testing locations, evaluation methods and results for the state extension trials are documented at their web sites (Washington: http://variety.wsu.edu/; accessed 21 Aug. 2012), (Oregon: http://cropandsoil.oregonstate.edu/wheat/state_performance_data.htm; accessed 21 Aug. 2012), (Idaho: http://www .extension.uidaho.edu/cereals/nidaho/; accessed 21 Aug. 2012) Data from the 2006–2010 Washington State Soft Winter Wheat Extension Trials (96 location-years) was used to compare Cara with the major club and soft white winter wheat cultivars in production in the region. Environments were grouped and analyzed within four annual precipitation zones plus one irrigated location. Each zone comprised four to six environments in each year, except for the irrigated location. Because of unequal variances across environments, data were analyzed using weighted mixed models, with the weight being equal to the inverse of the error variance for a trait within an environment. In these models, environment, genotype, and their interactions were considered fixed, and blocks within environments were considered random. Least squares means were calculated for genotypes within precipitation zones and significant differences among genotypes determined using Fisher’s protected F-test. In 2010 a severe stripe rust epidemic affected plant growth and grain yield, so the data were also analyzed as above but with environments grouped based on the effect of stripe rust on results rather than by precipitation zones.

The coleoptile length of Cara was evaluated based on 15 plants per cultivar and two replications per trial in 2009 and 2011. The experiments were conducted according to Hazkima et al. (2000) and analyzed using mixed models with year and cultivar considered fixed and replication and plant considered random. Cold tolerance was evaluated in artificial freezing trials conducted at the WSU Wheat Plant Growth center in two replications of 20 plants per replication for each of 3 yr from 2008 to 2010 as described in Skinner and Bellinger (2011). Logistic regression was performed using the LOGISTIC procedure of SAS/STAT software (SAS Institute, 2011) to compare the survival of Cara with that of the check cultivars Stephens (CItr 17596; sensitive) and Eltan (PI 536994; resistant). The conditions of each soil cell were recorded with temperature-monitoring probes and included in the analysis as covariates in the logistic regression. Confidence intervals of the ratios of odds of survival of Cara and other cultivars against Stephens and against Eltan were generated.

Resistance to P. striiformis races Pst37, Pst45, Pst100, Pst114, and Pst127 was evaluated on seedlings in the greenhouse as infection type (IT) on a scale of 1 to 9, where 1 = resistant and 9 = susceptible. Screening conditions and the rating system were as described in Line and Qayoum (1992). During the breeding process, adult-plant resistance to stripe rust was evaluated in field trials that had been artificially inoculated with Pst78 or Pst100. A small amount of spores was mixed with talc or flour and applied using a hand-held crop duster

in the spring as soon as the minimum night temperature rose above 0°C. No attempt was made to control natural infection, so actual populations of stripe rust in the field included both inoculated and naturally occurring races. Stripe rust was evaluated at Mt. Vernon and at Pullman WA under natural infection from 2004 through 2010. Prevalent races of stripe rust occurring in Pullman were Pst100, Pst116, and Pst127 (Chen, 2007; Chen et al., 2010). Ratings were recorded as IT and as severity (percentage of plot infected).

Resistance to lodging caused by strawbreaker foot rot was evaluated from 2006 to 2010, in headrows at Pullman WA with three replications. For these trials, infested oats were applied in the fall, 3 wk after planting. The space between rows was cultivated in the spring to cause root damage. Lodging was rated in July, after senescence. The rating scale in these plots was 1 = standing to 5 = flat. Resistance was also evaluated as described in Wetzel et al. (2009, 2010b).

Resistance to Cephalosporium gramineum was evaluated in inoculated field trials as described by Wetzel et al. (2010a). Resistance to powdery mildew was rated on a scale of 1 to 10 (Lipps and Madden, 1989) at St John, WA in 2005 and 2006 and at Moses Lake, WA in 2005.

The end use quality of Cara was evaluated by the USDA-ARS Western Wheat Quality Laboratory on grain samples from a total of 33 nurseries grown over 8 yr. Quality analyses for milling, baking, and product tests as listed in Table 6 were performed according to AACC international protocols (AACC International, 2008). Quality data for Cara were compared with Bruehl and Chukar as checks using t tests paired over environments. The actual number of environments (n) included in each analysis depended on whether the check cultivars were included in individual trials.

Cara was assayed with several molecular markers linked to resistance genes including Xbarc8 for Yr15, csLv34 for Lr34/Yr18, the Ventriup-LN2 marker for Yr17/Lr37/Sr38, the ORW1 marker for Pch1 and Xwmc525 for Pch2. Cara was also assayed with the UMN19 and UMN25 markers for alleles at glutenin loci and the specific markers for the wild-type and mutant alleles at Rht-D1 and Rht-B1 (Ellis et al., 2002; Helguera et al., 2003; Lagudah et al., 2006; Leonard et al., 2008; Liu et al., 2008). For additional details see Marker Assisted Selection in Wheat (http://maswheat .ucdavis.edu/protocols/index.htm; accessed 21 Aug. 2012). Marker assays were performed by the USDA-ARS Western Regional Small Grains Genotyping laboratory.

Characteristics and DiscussionAgronomic Characteristics

The coleoptile length of Cara is moderate compared with standard-height and other semidwarf checks (Table 1), indi-cating that Cara is better suited to planting with a standard grain drill rather than a deep furrow drill. Cara has the Rht-B1a wild-type and Rht-D1b mutant dwarfing allele derived from Norin-10. Cara has winter hardiness equal to that of other widely grown club wheat cultivars Bruehl, Chukar, and Coda, but it was not as good as the soft white wheat cul-tivars Eltan or Masami (PI 634715). Cara was significantly


In the Washington statewide variety trials from 2006 to 2010 (Table 3), the heading dates of Cara and Coda were same as those of the latest-heading cultivars, Bruehl, Chukar, and Eltan, in the dry environments and in the irrigated environment. Those environments had more locations that were warmer during the spring, and heading occurred over fewer days than in other precipitation zones. The heading date of Cara and Coda was intermediate to other cultivars when annual precipitation was between 30 and 60 cm. The plant height of Cara was less than that of most other cultivars in all precipitation zones. The grain protein concentration of Cara was intermediate to other cultivars. All cultivars had acceptable levels of grain protein for soft wheat produced in the PNW. The higher grain proteins in the irrigated environment may have been due to residual nitrogen in the cropping system or to terminal heat stress that occurred in that environment. The grain volume weight of Cara was less than that of the soft wheat cultivar checks in all environments but equal to that of the club wheat cultivars Bruehl and Chukar. Club wheat cultivars traditionally have a lower grain volume weight than other soft wheat classes and the U.S. grain grading system allows no. 1 grade club wheat to be up to 2.5 kg hL-1

less than other soft wheat classes (USDA 2006; U.S. Wheat Associates, Buyers Guide: http://www.uswheat.org/buyersGuide/wheatClasses; accessed 21 Aug. 2012). Grain yields of Cara were among the highest compared with those of other club and soft white cultivars in the 40- to 50-cm annual precipitation zone and were superior to those of the club wheat cultivars Bruehl, Chukar, and Coda in the 30- to 40-cm zone. Bruehl, Eltan, and Xerpha were superior to Cara in the lowest precipitation zone and ORCF102 and Xerpha were superior to Cara in the high-precipitation and irrigated zones.

The target growing area of Cara has experienced major epidemics of stripe rust since 2009. When data for 2006–2010 Washington Soft White Winter Wheat Trials were analyzed based on the effect of stripe rust on grain yields, Cara showed a clear advantage when stripe rust was a problem (Table 4). Because of its resistance, Cara has ranked among the top 10 genotypes for grain yield across all sites in the 2010 and 2011 Oregon Soft Winter Elite Yield Trials (total 23 site-years) It was among the top five cultivars at intermediate (20–55 cm) rainfall locations in the Washington Soft White Winter Wheat Trials in 2010 and 2011.

(p < 0.05) more tolerant to freezing than the susceptible check, Stephens, and significantly less tolerant than the resistant check, Eltan (Table 2).

In USDA-ARS plot trials from 2007 to 2009, Cara had a heading date the same as that of the club wheat checks, Chukar and Coda. In comparison with soft white wheat check cultivars, the heading date of Cara was also the same as that of Eltan, ‘Finch’ (PI 628640), and ‘Xerpha’ (PI 645605) but significantly later than ‘Brundage 96’ (PI 631486) (p < 0.02) and ‘Tubbs 06’ (p < 0.001). The height of Cara was equal to that of Chukar and Brundage 96 and shorter than that of the other cultivars (p < 0.01). The test weight of Cara was significantly (p < 0.001) lower than that of Coda, Eltan, Finch, Tubbs 06, and Xerpha but not different than that of the club check, Chukar. The grain yield of Cara was less than that of Tubbs 06 (p < 0.07) but equal to that of the other club and soft white cultivars.

Table 1. Coleoptile lengths, based on four trials, averaged across 15 seedlings.

Cultivar Length Standard errormm

Edwin 117.8 1.71 a†

Moro 114.7 1.72 a

Bruehl 87.3 1.69 b

Chukar 86.4 1.97 b

Eltan 83.6 1.95 bc

Xerpha 81.7 1.69 c

Cara 80.8 1.69 cd

Tubbs 06 76.8 1.95 def

Hiller 76.7 1.71 ef

Coda 73.7 1.95 f†Values followed by the same letter are not significantly different (p < 0.05).

Table 2. Survival comparisons among entries in the 2010 Washington State Extension Soft Wheat Variety Trial after freezing to −13°C.†

Worse than Stephens‡

Equal to Stephens

Better than Stephens, worse than Eltan

Equal to Eltan

Better than Eltan

Goetze Finch AP 700 CL CDC Ptarmigan

Skiles Lambert AP Legacy

Legion Bitterroot

ORCF-102 Bruehl

Rod Brundage 96

Simon Cara

WB-528 Chukar

Coda

George

Madsen

Masami

ORCF-103

RJames

Salute

Tubbs 06

Xerpha†Logistic regression was performed using the LOGISTIC procedure of SAS/STAT software (SAS Institute, 2011) to compare survival of entries to the check cultivars ‘Stephens’ (CItr17596) (sensitive) and ‘Eltan’ (PI536994) (resistant).

‡All entries in the same column are statistically similar. Entries in different columns are statistically different at p < 0.05.


Tab

le 3

. Ag

rono

mic

cha

ract

eris

tics

of

Car

a vs

. oth

er c

heck

s in

Was

hing

ton

Stat

e U

nive

rsit

y C

erea

l Var

iety

Tes

ting

Tri

als,

20

06

–201

0.†

Ann

ual p

reci

pit

atio

n‡

Cul

tiva

r<

30 c

m30

–40

cm40

–50

cm>

50 c

mIr

rig

ated

Gra

in v

olu

me

wei

ght

(kg

hL−

1 )B

rueh

l75

.7 d

73.0

f74

.3 i

73.7

f72

.8 f

Bru

ndag

e 96

75.9

d75

.0 c

76.1

ef

74.9

e75

.9 b

Car

a74

.6 e

73.7

e75

.6 g

73.9

f74

.1 e

Chu

kar

74.8

e74

.0 e

75.9

fg74

.5 e

74

.8 c

de

Co

da

78.4

a78

.0 a

79.1

a

78.3

a77

.2 a

Elt

an77

.1 b

75.3

c76

.8 d

76.1

c75

.1 b

cdM

adse

n76

.5 c

75.8

c77

.1 c

75.6

d75

.7 b

cO

RC

F-10

277

.2 b

76.5

b77

.8 b

76.7

b77

.4 a

Tub

bs

0675

.6 d

74.5

d75

.2 h

74.7

e74

.6 d

eX

erp

ha76

.6 c

75.2

c76

.3 e

75.8

cd

75.4

bcd

SE0.

140.

130.

110.

150.

35G

rain

yie

ld (k

g h

a−1 )

Bru

ehl

3232

cd

6644

d75

05 c

7937

bcd

10,2

84 d

eB

rund

age

9631

14 d

6876

bc

7714

bc

7983

bc

10,6

47 b

cdC

ara

3093

d66

63 c

7915

ab

7548

e99

52 e

fC

huka

r32

71 b

c65

22 d

8049

a78

32 c

d96

37 f

Co

da

3192

c65

22 d

7646

c79

11 b

cd95

03 f

Elt

an33

96 b

6621

d72

24 d

8047

bc

9901

ef

Mad

sen

3025

e64

81 e

7538

c77

12 d

e10

,524

cd

OR

CF-

102

3230

cd

6925

b80

77 a

8663

a11

,032

ab

Tub

bs

0631

94 c

d67

56 b

cd76

50 c

8130

b10

,721

bcd

Xer

pha

3569

a73

82 a

8046

a86

82 a

11,2

27 a

SE66

–71¶

9488

100

201

† Dat

a w

ere

anal

yzed

usi

ng m

ixed

mo

del

s w

ith

all f

acto

rs c

ons

ider

ed fi

xed

. Mo

del

s w

ere

wei

ght

ed b

y th

e re

cip

roca

l of t

he e

rro

r va

rian

ce w

ithi

n an

env

iro

nmen

t to

ad

just

for

vari

ance

het

ero

gen

eity

.‡ L

east

sq

uare

s m

eans

ove

r 25

, 23,

23,

29,

and

5 e

nvir

onm

ents

for

< 3

0, 3

0–4

0, 4

0–5

0, >

50,

and

irri

gat

ed

pre

cip

itat

ion

zone

s, r

esp

ecti

vely

.§ M

eans

follo

wed

by

the

sam

e le

tter

are

no

t si

gni

fican

tly

dif

fere

nt a

t a

= p

rob

. < 0

.05.

¶St

and

ard

err

ors

for

gra

in y

ield

in t

his

pre

cip

itat

ion

zone

var

ied

am

ong

cul

tiva

rs d

ue t

o lo

ss o

f plo

ts in

so

me

envi

ronm

ents

.

Ann

ual p

reci

pit

atio

n‡

Cul

tiva

r<

30 c

m30

–40

cm40

–50

cm>

50 c

mIr

rig

ated

Hea

din

g (d

ays

fro

m 1

Jan

uary

)B

rueh

l15

2.0

a§16

0.3

a15

9.5

b16

7.1

a15

2.6

aB

rund

age

9614

8.3

d15

5.8

g15

5.4

g16

3.0

e14

8.6

dC

ara

152.

1 d

158.

9 c

157.

9 d

165.

3 c

152.

3 ab

Chu

kar

152.

2 d

159.

3 b

158.

4 c

166.

0 b

152.

6 a

Co

da

151.

8 d

159.

0 c

158.

4 c

165.

2 c

152.

4 ab

Elt

an15

2.0

d16

0.2

a16

0.0

a16

7.1

a15

2.4

aM

adse

n15

0.9

b15

8.5

d15

7.9

d16

5.4

c15

2.0

abO

RC

F-10

214

9.2

c15

6.5

f15

6.2

e16

3.1

e14

9.2

dTu

bb

s 06

149.

3 c

156.

5 f

155.

8 f

163.

5 d

150.

5 c

Xer

pha

150.

9 b

158.

4 e

157.

8 d

165.

3 c

151.

6 b

SE0.

130.

110.

130.

140.

32P

lant

hei

ght

(cm

)B

rueh

l78

.7 a

93.7

bc

97.4

cd

100.

5 b

c95

.7 d

Bru

ndag

e 96

70.8

e87

.9 e

90.9

g93

.2 f

91.9

eC

ara

67.7

f85

.2 f

89.8

f90

.3 g

93.8

de

Chu

kar

71.6

e89

.3 d

94.9

e96

.0 e

98.4

bc

Co

da

75.6

c94

.4 b

99.3

ab

101.

6 ab

100.

0 ab

Elt

an76

.8 b

c94

.1 b

96.6

d10

0.0

c98

.9 b

Mad

sen

74.3

d89

.9 d

93.3

f95

.7 e

96.3

cd

OR

CF-

102

77.3

b96

.1 a

98.4

ab

c10

2.0

a98

.4 b

cTu

bb

s 06

77.5

ab

96.0

a10

0.0

a10

1.1

abc

102.

2 a

Xer

pha

77.5

ab

92.6

c96

.6 d

97.4

d10

0.5

abSE

0.50

0.54

0.52

0.49

1.04

Gra

in p

rote

in c

onc

entr

atio

n (%

)B

rueh

l12

.1 a

b11

.2 a

11.7

ab

10.8

bc

12.9

bB

rund

age

9611

.8 c

d10

.8 b

11.5

cd

10.6

cd

12.4

cC

ara

11.9

bc

10.8

b11

.2 e

10.9

ab

13.4

aC

huka

r11

.6 d

10.7

bc

11.0

f10

.6 c

d13

.4 a

Co

da

12.1

ab

11.0

ab

11.8

ab

10.9

ab

13.4

aE

ltan

11.6

d10

.8 b

11.4

cd

e10

.4 e

12.7

bc

Mad

sen

12.2

a11

.2 a

11.9

a11

.0 a

12.6

cO

RC

F-10

211

.9 b

c10

.9 b

11.6

bc

10.7

cd

12.3

dTu

bb

s 06

11.7

cd

10.7

bc

11.3

de

10.4

de

12.3

dX

erp

ha11

.3 e

10.5

c11

.5 c

d10

.5 d

e12

.4 c

dSE

0.10

0.11

0.08

0.09

0.15


Resistance to Plant PathogensCara was highly resistant (IT of 2) at seedling stages to all tested races of stripe rust in the greenhouse compared with an IT of 8 for the susceptible check, WA7821. In the field at flowering to soft dough stages, since 2003, Cara has con-sistently exhibited a resistant reaction (IT of 2 and sever-ity of 5–10%) when the susceptible check, WA7821, was rated with an IT of 8 and severity of 60 to 100% (data for 2010 and 2011 is in Table 5). Cara has the same allele at the Ventriup marker as Madsen, indicating that it probably has Yr17. Current races of stripe rust that are prevalent in the PNW, however, are virulent to Yr17, indicating that Cara has other genes for resistance possibly derived from PI 178383 and Tres. The stripe rust races currently prevalent in the PNW, in combination, are virulent to all resistance genes currently identified except for Yr5 and Yr15. The sources of those genes are not present in the pedigree of Cara. Research to identify the combination of genes responsible for stripe rust resistance in Cara are currently underway.

Cara is resistant to strawbreaker foot rot and has the Pch1 gene for resistance to strawbreaker foot rot derived from Madsen. Cara also possesses the same allele at Xwmc525 as ‘Cappelle Desprez’ (PI 518787), which is associated with resistance derived from the Pch2 gene (Chapman et al., 2008). Cappelle Desprez was not a direct

Table 4. Grain yield of Cara compared with checks from WSU Extension Cereal Variety Trials, by effect of stripe rust on trial environment.†

Cultivar Affected‡ Not affected————————— kg ha−1 —————————

Bruehl 7246 ef§ 6239 de

Brundage 96 7640 cd 6279 d

Cara 8102 a 6029 g

Chukar 7939 ab 6152 ef

Coda 7487 de 6111 fg

Eltan 6680 h 6321 cd

Madsen 7404 de 6069 g

ORCF-102 7764 bc 6595 b

Tubbs 06 7082 fg 6398 c

Xerpha 6845 gh 7008 a

SE 95.1 35.8†Data were analyzed using mixed models with all factors considered fixed. Models were weighted by the reciprocal of the error variance within an environment to adjust for variance heterogeneity.

‡Least squares means over 17 and 79 environments for stripe rust affected and unaffected, respectively.

§Means followed by the same letter are not significantly different at p < 0.05.

Table 5. Infection type (IT) and severity (% infection) for stripe rust based on natural infection of major soft wheat cultivars grown in Washington in 2010 and 2011.

Location†

Spillman Plant Pathology Whitlow Mt Vernon, WA Walla Walla, WA Lind, WARating date 2010: 22 June 23 June 24 June 14 Apr. 24 May 10 June 17 June

Rating date 2011: 23 June 22 June 24 June 10 May 4 June 10 June 16 JuneCultivar Year IT % IT % IT % IT % IT % IT % IT % Summary

Bruehl 2010 2 5 2 2 2 2 2 5 2 5 2 1 2 2 R

2011 2 5 2 5 2 5 2 5 3 5 2 5 3 2

Brundage 96 2010 5 30 3 30 8 20 5 20 2 10 3 20 3 5 MR

2011 5 50 2 20 3 20 2 10 3 10 3 20 5 30

Cara 2010 2 1 2 2 2 2 2 2 1 1 2 1 2 2 R

2011 2 2 2 1 2 2 2 1 3 5 2 5 3 2

Chukar 2010 2 1 2 2 2 5 2 5 2 1 2 1 2 2 R

2011 2 5 2 1 2 2 2 5 2 1 2 5 3 5

Eltan 2010 3 20 5 20 2 20 5 20 2 5 3 20 2 10 MR

2011 2 20 2 10 3 10 2 5 2 10 2 10 5 40

Madsen 2010 2 1 0 0 2 5 5 20 3 20 2 1 2 2 R

2011 2 5 2 2 2 5 2 5 2 5 2 5 3 5

ORCF-102 2010 3 30 5 10 8 30 8 60 3 30 8 20 2 2 MR

2011 5 50 5 20 8 50 3 10 3 15 5/2 20 5 30

Stephens 2010 3 40 5 30 5 10 3 10 2 30 3 30 2 5 MR

2011 5 60 2 40 3 30 3 10 2 10 3 50 5 20

Tubbs 06 2010 8 40 8 50 8 40 8 40 5 30 8 90 8 15 S

2011 8 90 8 60 8 80 8 20 8 30 3 40 8 40

XERPHA 2010 8 50 8 30 8 40 8 50 3 30 5 50 8 10 MS

2011 7 80 5 40 7 80 8 60 5 20 5 10 5 60

WA7821(susceptible check) 2010 8 80 8 80 8 80 8 60 8 90 8 100 8 60 S

2011 8 100 8 90 8 90 8 80 8 80 8 70 8 100†Spillman, Plant Pathology, and Whitlow farms are near Pullman, WA.


parent of Cara, but it is in the pedigree of Moisson. Mean lodging for Cara across 2 yr (2010 and 2011) in the presence of severe foot rot was 1.08, similar to the resistant check Madsen (1.35) and significantly (p < 0.001) less than the susceptible cultivars Xerpha (3.42), Bruehl (4.75), and Eltan (4.86). In plot trials conducted by Wetzel et al. in 2008 and 2009, Cara was rated as having a similar disease incidence, severity, and index as Madsen and was significantly (p < 0.05) better than the susceptible cultivars Bruehl and Eltan.

Cara has resistance to field races of powdery mildew in Washington and was rated 2–3 on the 1–9 scale of Lipps and Madden (1989) where susceptible entries were rated 7–9 at Moses Lake, Pullman, and St. John, WA in 2005. Powdery mildew has been observed on Cara in the greenhouses at WSU, however. Powdery mildew is not a frequent disease in the dryland production region of Washington, and races present in PNW field and greenhouses are unknown. Cara was rated moderately susceptible to Cephalosporium gramineum (Wetzel et al., 2010a).

End-Use QualityThe milling, baking, and product quality of Cara were as good as those of the best club wheat check cultivars with high break flour and cake volumes (Table 6). Cara was sig-nificantly better than Bruehl and equal to Chukar for grain volume weight. Cara had a significantly smaller kernel size than Bruel, but it was equal to that of Chukar. Grain pro-tein content and flour protein content of Cara were equal to those of Bruehl but the flour protein content of Cara was slightly higher than that of Chukar. The milling quality of Cara is typical of club wheat cultivars. The flour yield and break flour yield of Cara were significantly better than those of Bruehl and equal to those of Chukar. Flour ash content was the same for all three cultivars, and the overall milling score for Cara was better than for Bruehl and equal to Chukar. The single-kernel hardness value of Cara was higher than Bruehl and equal to Chukar. Although single-kernel hardness is a predictor of grain texture, it measures crush-ing resistance rather than endosperm separation and is not 100% correlated with break flour yield. The mixograph water absorption was low and equal for all three cultivars. The product tests indicated that Cara has significantly bet-ter cookie diameter than Bruehl, but not as good as Chukar, and the cake volumes were equal for all three cultivars. Cara has either the Ax1 or AxNull subunit at Glu-A1 based on the UMN19 STS marker. The UMN25 marker indicated that Cara has the Dx5/Dy10 subunits at Glu-D1. Cara is an awnless,

white-chaffed, club wheat with white kernels. Samples of grain sent for grading to the Federal Grain Inspection Ser-vice were rated as White Club or Western White wheat in both 2005 and 2006.

In summary, Cara is agronomically competitive with other soft wheat cultivars in the PNW and has excellent resistance to stripe rust and strawbreaker foot rot combined with excellent club wheat end-use quality. The weaknesses are similar to those of other club wheat cultivars in the PNW; it is not as winter hardy, nor does it have grain

Table 6. End-use quality of Cara club wheat compared with the two Pacific Northwest club wheat cultivars occupying the most acreage in Washington since 2002, compared using balanced paired t tests.

Trait means

Cultivar Cara Check N LSD p valueGrain volume weight (kg hL−1)

Bruehl 75.25 74.38 21 0.68 0.02

Chukar 77.25 76.63 15 0.79 0.09

Grain protein (%)

Bruehl 10.0 10.0 21 0.55 0.80

Chukar 10.4 9.9 15 0.48 0.08

Single-kernel hardness (0–100 index)

Bruehl 35.5 30.6 21 1.66 0.001

Chukar 35.3 33.4 15 2.05 0.07

Kernel weight (g 1000 kernels−1)

Bruehl 31.6 36.2 21 1.33 0.001

Chukar 32.7 32.6 15 0.99 0.73

Flour yield (%)

Bruehl 71.3 70.1 21 0.46 0.001

Chukar 71.2 70.7 15 0.66 0.13

Break flour yield (%)

Bruehl 53.6 50.9 21 0.53 0.001

Chukar 51.7 52.0 15 0.59 0.28

Flour ash (%)

Bruehl 0.38 0.396 19 0.02 0.08

Chukar 0.35 0.356 15 0.02 0.55

Milling score (0–100 index)

Bruehl 87.8 85.3 19 1.4 0.001

Chukar 89.4 88.5 15 1.08 0.09

Flour protein (%)

Bruehl 8.5 8.6 21 0.55 0.80

Chukar 8.6 8.2 15 0.41 0.04

Mixograph water absorption (mL)

Bruehl 51.7 51.5 21 0.93 0.73

Chukar 50.5 49.4 9 2.01 0.27

Cookie diameter (cm)

Bruehl 9.8 9.6 21 0.06 0.001

Chukar 9.5 9.7 15 0.1 0.01

Cake volume (cc)

Bruehl 1286 1286 21 25 0.98

Chukar 1255 1289 11 55 0.20


Kidwell, K.K., V.L. DeMacon, G.B. Shelton, J.W. Burns, B.P. Carter, C.F. Morris, and X.M. Chen. 2004. Registration of ‘Eden’ wheat. Crop Sci. 44(5):1870–1871. doi:10.2135/cropsci2004.1870

Lagudah, E.S., H. McFadden, R.P. Singh, J. Huerta-Espino, H.S. Bariana, and W. Spielmeyer. 2006. Molecular genetic characterization of the Lr34/Yr18 slow rusting resistance gene region in wheat. Theor. Appl. Genet. 114:21–30. doi:10.1007/s00122-006-0406-z

Leonard, J.M., C.J.W. Watson, A.H. Carter, J.L. Hansen, R.S. Zemetra, D.K. Santra, K.G. Campbell, and O. Riera-Lizarazu. 2008. Identification of a candidate gene for the wheat endopeptidase Ep-D1 locus and two other STS markers linked to the eyespot resistance gene Pch1. Theor. Appl. Genet. 116:261–270. doi:10.1007/s00122-007-0664-4

Lin, P.-Y., and S. Czuchajowska. 1997. Starch properties and stability of club and soft white winter wheats from the Pacific Northwest of the United States. Cereal Chem. 74:639–646. doi:10.1094/CCHEM.1997.74.5.639

Line, R., and A. Qayoum. 1992. Virulence, aggressiveness, evolution, and distribution of Puccinia striiformis (the cause of stripe rust in wheat) in North America 1960–1987.USDA Tech. Bull. 1788, USDA, Washington, DC.

Lipps, P., and L. Madden. 1989. Assessment of methods of determining powdery mildew severity in relation to grain yield of winter wheat cultivars in Ohio. Phytopathology 79:462–470. doi:10.1094/Phyto -79-462

Liu, S., S. Chao, and J.A. Anderson. 2008. New DNA markers for high molecular weight glutenin subunits in wheat. Theor. Appl. Genet. 118:177–183. doi:10.1007/s00122-008-0886-0

Morris, C.F., K.G. Campbell, and G.E. King. 2005. Kernel texture differences among US Soft wheat cultivars. J. Sci. Food Agric. 85:1959–1965. doi:10.1002/jsfa.2202

Peterson, C.J., C.F. Morris, R.F. Line, E. Donaldson, S.S. Jones, and R.E. Allan. 1999. Registration of ‘Hiller’ wheat. Crop Sci. 39(5):1531.

Peterson, C.J., O.A. Vogel, D.W. George, and G.L. Rubenthaler. 1977. Registration of Barbee wheat. Crop Sci. 17(4):675.

Rohde, C.R. 1966. Registration of Moro wheat. Crop Sci. 6(5):502. SAS Institute. 2011. SAS/STAT 9.3 user’s guide. SAS Institute, Cary, NC.Skinner, D.Z., and B.S. Bellinger. 2011. Differential response of wheat

cultivars to components of the freezing process in saturated soil. Plant Soil 51:69–74.

USDA. 2006. Subpart M. United States standards for wheat. U.S. Standards for grain. http://www.gipsa.usda.gov/fgis/standards/810wheat.pdf (accessed 21 Aug. 2012).

USDA National Agricultural Statistics Service (NASS). 2011. Washington’s wheat variety–2011 crop. http://www.nass.usda .gov/Statistics_by_State/Washington/Publications/Small_Grains/index.asp (accessed 21 Aug. 2012).

Wetzel, H.D., III, J.C. Evans, K.E. Klos, and T.D. Murray. 2010a. Reaction of winter wheat cultivars and breeding lines to Cephalosporium stripe, 2009. Plant Dis. Manage. Rep. 4:CF027. http://www.plantmanagementnetwork.org/pub/trial/pdmr/volume4/abstracts/cf27.asp (accessed 21 Aug. 2012).

Wetzel, H.C., III, J.C. Evans, and T.D. Murray. 2010b. Reaction of winter wheat cultivars and breeding lines to eyespot, 2009. Plant Dis. Manage. Rep. 4:CF028. http://www.plantmanagementnetwork.org/pub/trial/pdmr/volume4/abstracts/cf28.asp (accessed 21 Aug. 2012).

Wetzel, H.C., III, K.E. Klos, and T.D. Murray. 2009. Reaction of winter wheat cultivars and breeding lines to eyespot, 2008. Plant Dis. Manage. Rep. 3:CF041. http://www.plantmanagementnetwork.org/pub/trial/pdmr/volume3/abstracts/cf41.asp (accessed 21 Aug. 2012).

Zadoks, J.C., T.T. Chang, and C.F. Konzak. 1974. A Decimal Code for the Growth Stages of Cereals. Weed Res. 14:415–421. doi:10.1111/j.1365-3180.1974.tb01084.x

Zwer, P.K., A. Sombrero, R.W. Rickman, and B. Klepper. 1995. Club and common wheat yield component and spike development in the Pacific-Northwest. Crop Sci. 35:1590–1597. doi:10.2135/cropsci1995.0011183X003500060012x

volume weight equal to recently released soft white winter wheat cultivars.

AvailabilityFifteen hundred heads were selected from a purification plot grown at Spillman Farm in 2005 and given to the Washington State Crop Improvement Association and increased as F14 headrows in 2006. Because of off-types in the 2007 breeder seed production, 1500 heads were again selected in 2007 and grown as F16 headrows in 2008 for a second breeder seed production. Cara is sold as Founda-tion, Registered, and Certified seed classes. Small amounts (5 g) of seed are available from the corresponding author for research purposes. Seed of Cara has been deposited in the National Small Grains Collection and the National Plant Germplasm system, where it will be available for distribu-tion by NPGS 5 yr after the date of publication.

ReferencesAACC International. 2008. Approved methods of the AACCI, 11th ed.

AACCI, St. Paul, MN. http://www.aaccnet.org/approvedmethods/default.aspx (accessed 21 Aug. 2012).

Allan, R.E., C.J. Peterson, G.L. Rubenthaler, R.F. Line, and K.J. Morrison. 1986. Registration of ‘Tres’ wheat. Crop Sci. 26(1):203. doi:10.2135/cropsci1986.0011183X002600010062x

Allan, R.E., C.J. Peterson, G.L. Rubenthaler, R.F. Line, and D.E. Roberts. 1989. Registration of ‘Madsen’ wheat. Crop Sci. 29(6):1575–1576. doi:10.2135/cropsci1989.0011183X002900060068x

Allan, R.E., C.F. Morris, R.F. Line, J.A. Anderson, M.K. Walker-Simmons, and E. Donaldson. 2000. Registration of ‘Coda’ club wheat. Crop Sci. 40(2):578.

Chapman, N.H., C. Burt, H. Dong, and P. Nicholson. 2008. The development of PCR-based markers for the selection of eyespot resistance genes Pch1 and Pch2. Theor. Appl. Genet. 117:425–433. doi:10.1007/s00122-008-0786-3

Chen, X. 2007. Challenges and solutions for stripe rust control in the United States. Aust. J. Agric. Res. 58:648–655. doi:10.1071/AR07045

Chen, X., L. Penman, A. Wan, and P. Cheng. 2010. Virulence races of Puccinia striiformis f. sp. tritici in 2006 and 2007 and development of wheat stripe rust and distributions, dynamics, and evolutionary relationships of races from 2000 to 2007 in the United States. Can. J. Plant Pathol. 32:315–333. doi:10.1080/07060661.2010.499271

Ellis, M.H., W. Spielmeyer, K.R. Gale, G.J. Rebetzke, and R.A. Richards. 2002. “Perfect” markers for the Rht-B1b and Rht-D1b dwarfing genes in wheat. Theor. Appl. Genet. 105:1038–1042. doi:10.1007/s00122-002-1048-4

Garland Campbell, K.A., R.E. Allan, J. Anderson, J.A. Pritchett, L.M. Little, C.F. Morris, R.F. Line, X. Chen, M.K. Walker-Simmons, B.P. Carter, J.W. Burns, S.S. Jones, and P.E. Reisenauer. 2005. Registration of ‘Chukar’ Wheat. Crop Sci. 45:1657–1658. doi:10.2135/cropsci2004.0342

Helguera, M., I.A. Khan, J. Kolmer, D. Lijavetzky, Z. Li, and J. Dubcovsky. 2003. PCR Assays for the Lr37-Yr17-Sr38 cluster of rust resistance genes and their use to develop isogenic hard red spring wheat lines. Crop Sci. 43:1839–1847. doi:10.2135/cropsci2003.1839

Hakizimana, F., S.D. Haley, and E.B. Turnipseed (2000). Repeatability and genotype × environment interaction of coleoptile length measurements in winter wheat. Crop Sci. 40: 1233–1237.

Johnson, E.B., J.V.J. Nalam, R.S. Zemetra, and O. Riera-Lizarazu. 2008. Mapping the compactum locus in wheat (Triticum aestivum L.) and its relationship to other spike morphology genes of the Triticeae. Euphytica 163:193–201. doi:10.1007/s10681-007-9628-7

Jones, S.S., E. Donaldson, S.R. Lyon, C.F. Morris, R.F. Line, and R. Hoffman. 2000. Registration of ‘Edwin’ wheat. Crop Sci. 40(4):1198.

Jones, S.S., T.D. Murray, S.R. Lyon, C.F. Morris, and R.F. Line. 2001. Registration of ‘Bruehl’ wheat. Crop Sci. 41(6):2006–2007. doi:10.2135/cropsci2001.2006

Registration of 'Cara' Soft White Winter Club Wheat

Documents

Transcript of Registration of 'Cara' Soft White Winter Club Wheat